WASHINGTON -- Twitter suspended a string of popular parody accounts lampooning Russian officials, sparking outrage among their followers and accusations that the social-media giant is bending to pressure from the Kremlin and its allies.

At least five English-language Twitter accounts needling the Russian government had been suspended as of May 31, though three of those were later reinstated after apparently appealing to the California-based company.

The popular @DarthPutinKGB account, however, remained suspended late on May 31. The Twitter feed mocks Russian President Vladimir Putin, playfully asserting that it is, in fact, the one being parodied by the Kremlin’s official Twitter feed.

Fans of the feed swiftly launched the hashtag campaign #NoTwitterGulagForDarthPutinKGB in support.

Three of the suspended accounts that were later unfrozen were @RusEmbassyNot, a parody of Russian diplomacy efforts; @SovietSergey, which tweaks Russian Foreign Minister Sergei Lavrov; @AmbYakovenkoNot, which lampoons Aleksandr Yakovenko, Russia’s ambassador to Britain.

The @Russia__Not account, which spoofs the Russian government, also remained suspended late on May 31.

Twitter spokesman Ian Plunkett declined to provide an explanation for the suspensions, saying in an e-mail that Twitter does “not comment on individual accounts for privacy and security reasons.” He referred RFE/RL to the California-based company’s rules for parody accounts.

Those rules say that the feed’s "bio" should explicitly state that it is not affiliated with the actual subject by using words like “parody” or “fake.” It also states that the account name “should not be the exact name of the account subject without some other distinguishing word, such as ‘not,’ 'fake,' or 'fan.'"

Both of these criteria should be fulfilled “in a way that would be understood by the intended audience,” the rules state.

Both Facebook and Twitter in recent years have removed posts and accounts critical of Russian authorities based on claims by Russian state media watchdog Roskomnadzor that they violate domestic laws. In some cases they have been restored shortly thereafter.

Yury Melnik, a spokesman for the Russian Embassy in Washington, said that they had no information about Russian officials asking Twitter to suspend the accounts.

"The embassy certainly did not get in touch with Twitter," he wrote in an e-mail to RFE/RL.

Kremlin opponents have also accused online “trolls” allegedly bankrolled by the Russian government of flooding both Twitter and Facebook with complaints in order to smother criticism of Moscow on the Internet. Amid the government’s steady tightening of its control over the media, the Internet has remained an important forum in Russia for dissent and uncensored debate.

But the man who says he runs two of the frozen accounts in question, aside from @DarthPutinKGB, told RFE/RL in an online chat that he believes the Russian Foreign Ministry may have been behind the Twitter suspensions.

He noted that the accounts were frozen shortly after Foreign Ministry spokeswoman Maria Zakharova complained on Facebook about a Euronews article that had embedded a tweet from @SovietSergey but later removed it.

The man declined to identify himself, though RFE/RL was able to corroborate his affiliation with the accounts through screenshots. He said he is directly responsible for two of the accounts, and collaborates on the other two.

Zakharova called the inclusion of the tweet in an article about an Italian region urging the national government to lift sanctions against Russia “disinformation” whose “possible goal is to promote a fake outlet.”

The Kremlin has repeatedly accused the Western media and governments of vilifying Putin and his government.

In the two days following Zakharova’s statement, Twitter suspended the @SovietSergey, @AmbYakovenkoNot, and @Russia_Not accounts, the author of the feeds told RFE/RL. The @RusEmbassyNot feed was suspended later.

He provided screenshots demonstrating his access to the four accounts:

“All of these are spoofs of either [Russian] diplomats or accounts run by them, or [Ministry of Foreign Affairs]," he said on condition of anonymity, citing concerns about “being targeted by Russia.” “There’s no troll campaign this time. I would have seen it on my notifications.”

In the case of the @SovietSergey account, Twitter notified him that if and when the suspension is lifted, he must remove the account name “Sergey Lavrov,” as well as the bio, which reads: “Not The Foreign Minister of the Russian Federation World Record Holder Of Denying Since 2008. Zakharova’s Boss. I Deny Being A Parody.”

The account was reactivated later on May 31 with the offending language deleted.

The suspension of @DarthPutinKGB, in particular, triggered a surge of condemnations from supporters of the account, which had attracted tens of thousands of followers with wry tweets like: "Don't believe anything the Kremlin doesn't first deny."

A blog maintained by the person behind @DarthPutinKGB wrote in a characteristically ironic post on May 31 that the Twitter suspension “is a Russophobic stab in the back by the accomplices of terrorists!”

“I will be back,” the blog post stated, punctuating the vow with an expletive.

The author of @DarthPutinKGB, meanwhile, told RFE/RL he started his parody feed in 2013 because he “didn’t believe anyone was properly parodying” Putin. He denied that the account violates Twitter’s rules on parody accounts, and said he expected to get a response to his request for the account to be unfrozen sometime this week.

“No sensible person could read my bio and think it is really the president of Russia. Anyone who can’t see that is being willfully stupid,” he said in an online chat.

“I wanted Darth in the user name as [Putin] is definitely from the dark side,” he said. “Plus, he’s KGB to his core. He’d deny it snows in Russia if [he] had to.”

He added that he had requested that Twitter unfreeze the account, and that he expected a decision this week.

One account that the author of the Lavrov parody feed said he was "95 percent" responsible for -- @Sputnik_Not, a parody of the state-run news agency -- remained active on May 31, and it took a direct swipe at Twitter's move: